The reproductive biology of the genusBaldellia (Alismataceae)

The reproductive biology of the genusBaldellia was studied under cultivation and in nature.B. ranunculoides subsp.repens is self-incompatible whereasB. ranunculoides subsp.ranunculoides andB. alpestris are self-compatible. Bud-pollination occurs in theB. alpestris. Self-incompatibility correlates with the production of fewer, larger, more showy flowers, higher pollen production (but reduced pollen grain size), fewer carpels and smaller seeds. Resource allocation to sexual reproduction is inversely correlated with the degree of vegetative growth.B. ranunculoides subsp.repens produces more ramets than subsp.ranunculoides, which reproduces mainly by seeds.B. alpestris is intermediate between the two since it is self-compatible, produces fewer, larger seed than subsp.ranunculoides and is also able to propagate by vegetative means.     

A new infraspecific arrangement of Teucrium thymifolium (Lamiaceae), an aggregate endemic to the Iberian Peninsula

In the framework of our studies on Lamiaceae from the eastern Iberian Peninsula, for the forthcoming fourth volume of Flora valentina, a new infraspecific arrangement is proposed here for the Iberian endemic Teucrium thymifolium. Besides the type subspecies, T. thymifolium subsp. thymifolium, which is widely distributed through the eastern Iberian Peninsula, two new nomenclatural combinations are established: (i) T. thymifolium subsp. fraternum (Pau) M.B. Crespo, M.A. Alonso & Mart.-Azorín (= T. terciae), for the thermophilous populations growing in SE Murcia province, and (ii) T. thymifolium subsp. hervieri (Briq. & Debeaux) M.B. Crespo, M.A. Alonso & Mart.-Azorín, for Subbaetic inland populations from where the provinces of Albacete, Jaén and Granada come into contact. Nomenclatural types and data on distribution and ecology are reported for all accepted subspecies, including the designation of a lectotype for T. thymifolium subsp. hervieri. Furthermore, relationships of the latter to the recently described T. moleromesae Sánchez Gómez et al. are briefly discussed, both taxa being treated here in synonymy.     

A study of Sarcocornia A.J. Scott (Chenopodiaceae) from Western Mediterranean Europe

The latest publications on Sarcocornia taxonomy and phylogeny recognize six taxa in this genus on the Iberian Peninsula: S. perennis, S. fruticosa, S. alpini, S. alpini subsp. carinata, S. hispanica, and S. pruinosa. The present study represents a comprehensive revision of the different taxa in the Sarcocornia genus present in Western Mediterranean Europe by means of morphological, micromorphological and phylogenetic internal transcribed spacer (ITS) analysis. Morphological and micromorphological data were studied from Sarcocornia samples from 113 populations in coastal salt marshes and inland salt pans in Portugal, France, Spain and Italy. Sixteen new ITS sequences were obtained from Mediterranean Sarcocornia species and analysed together with previous reported data. Published karyological, ecological and biogeographical data from Western Mediterranean Europe were also reviewed. The results indicate the presence of a new species, S. lagascae, found growing in coastal Mediterranean areas of the Iberian Peninsula. The species S. fruticosa was found to be absent from the Iberian territories.     

A new taxon in the genusMoehringia (Caryophyllaceae)

The new taxonMoehringia intricata subsp.giennensis, from the calcareous mountains of the south-eastern Iberian Peninsula is described. A comparative morphological study with regard to the most closely related taxa, contributes information of leaf anatomy, seed and pollen morphology, ecology and distribution.     

Morphological analysis,chorology and ecology of Juniperus navicularis Gand. in the Iberian Peninsula

Juniperus navicularis is an endemic taxon of Iberian Peninsula that develops on paleodunes of coastal ecosystems and whose distribution range is not well defined. In this paper, we present the results of the morphological analysis in populations attributed to J. navicularis of southern Spain. In order to assess the similarity between Portuguese and Spanish populations, the Mann–Whitney U test was performed, the same test was also applied between this taxon and Juniperus oxycedrus subsp. oxycedrus. In order to distinguish this species from other taxa of the Juniperus oxycedrus group (subsp. oxycedrus, subsp. badia, subsp. macrocarpa) in the southwestern Mediterranean region, a principal component analysis has been carried out. New data about the chorology and ecology of Spanish populations considered as a priority habitat in the Habitat Directive as well as number of individuals and conservation status are provided.     

Taxonomic revision of the genusPinguicula L. in the Iberian Peninsula

In this paper, the genusPinguicula is revised in the Iberian Peninsula. The taxa studied areP. vulgaris, P. nevadensis, P. grandiflora subsp.grandiflora, P. dertosensis, P. vallisneriifolia, P. longifolia subsp.longifolia, P. mundi, P. alpina andP. lusitanica. Five taxa are endemic to the area studied (P. nevadensis, P. dertosensis, P. vallisneriifolia, P. longifolia subsp.longifolia andP. mundi). Two of them are confined to restricted areas:P. nevadensis is endemic to the Sierra Nevada (S Spain) andP. longifolia subsp.longifolia is endemic to the central Pyrenees (N Spain). Taxonomical, nomenclatural, morphological, chromosomal, biogeographical and ecological data are reported for each taxon. A new combination is proposed:Pinguicula Sect.Longifolia (Casper) G. Blanca, M. Ruíz-Rejón etR. Zamora.     

First record of Latonia gigantea (Anura,Alytidae) from the Iberian Peninsula

The single extant species of the anuran genus Latonia lives in Israel, but in the fossil record the genus is known mainly from Europe, spanning from the Oligocene to the early Pleistocene. Here we describe new remains of Latonia from the early to late Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula), coming from the following localities: Sant Mamet (MN4), Sant Quirze and Trinxera del Ferrocarril (MN7+8), and Castell de Barberà, Can Poncic 1 and Can Llobateres 1 (MN9). Fossils from the late Aragonian and early Vallesian are attributed to Latonia gigantea mainly because of the morphology of the ornamentation that covers the maxillae. In turn, an ilium from Sant Mamet is not diagnostic at the specific level and is assigned only to the genus Latonia. The newly reported remains represent the first record of L. gigantea in the Iberian Peninsula, where Latonia was previously known by a single report of Latonia cf. ragei from Navarrete del Río (MN2) and remains from other localities unassigned to species. Moreover, the Vallès-Penedès remains represent one of the southernmost records of the species known thus far. The presence of Latonia in these localities confirms the humid and warm environment suggested by the recorded mammal fauna.     

Bryophyte flora of the Sierras de Filabres,Cabrera, Alhamilla and Cabo de Gata (Almería,S.E. Spain)

Abstract

The bryophyte flora of the Sierras de Filabres, Cabrera, Alhamilla and Cabo de Gata of Almería province (S.E. Spain) includes 280 taxa (236 mosses and 44 liverworts) of which 74 are new to the province of Almería, 31 to the southeastern part of the Iberian Peninsula and 4 to the Iberian Peninsula. Data about their biogeography and ecology are given.     

The genus Mantisalca (Asteraceae) in southwest Europe

The diversity of the genus Mantisalca in southwest Europe is the subject of much controversy. The present taxonomic revision of the genus on the Iberian Peninsula allows four species to be recognized, whose morphological variability is described and various synonyms are typified. A new species is described, M. cabezudoi Ruiz de Clavijo and Devesa, endemic to southeast Spain. The name Microlonchus spinulosus Rouy is reinstated for a species endemic to the central and east Iberian Peninsula, for which a new combination is made: Mantisalca spinulosa (Rouy) Ruiz de Clavijo and Devesa. A diagnostic key to the taxa is given.     

The Iberian Peninsula: ancient history of a hot spot of mite harvestmen (Arachnida: Opiliones: Cyphophthalmi: Sironidae) diversity

The Iberian Peninsula represents a hot spot of cyphophthalmid (mite harvestman) disparity, with four of the eight genera currently recognized in the family Sironidae represented in the region – a generic diversity and morphological disparity not found in any other region of the World so far. From these, two genera (Iberosiro and Odontosiro) are monotypic, and are restricted to the western side of the peninsula. Parasiro is restricted to the north‐east region, from the Catalonian Coastal Ranges and both sides of the Eastern Pyrenees, in areas where the annual rainfall surpasses 1000 mm, and mostly restricted to areas with Paleozoic and Variscan rocks, with other species of the genus extending to Corsica, Sardinia, and the Italian Peninsula. A second species of the genus Paramiopsalis, Paramiopsalis eduardoi sp. nov. from Fragas do Eume, is described here along with a re‐diagnosis of the genus. Paramiopsalis species, together with Odontosiro, inhabit the north‐west corner of the Iberian Peninsula, an area with some of the highest recorded annual rainfall, and with Paleozoic rocks from the Iberian Massif or Variscan granitoid rocks. A phylogenetic analysis of the members of the family Sironidae using four molecular markers, despite not including all of the Iberian genera, clearly shows the non‐monophyly of the Iberian Cyphophthalmi, indicating that the Iberian Peninsula is home to multiple ancient lineages of mite harvestmen. The two Paramiopsalis species form a sister clade to the Balkan genus Cyphophthalmus, whereas Parasiro constitutes the first lineage of the sironids represented.     

Four new species of Typhlocharis (baetica group) (Coleoptera: Carabidae: Anillini) from southwestern Iberian Peninsula with notes on their biogeographical and morphological implications

Four new species of Typhlocharis (Carabidae: Anillini) are described from the southwest of the Iberian Peninsula, assigned to the baetica species group. T. prima sp.n. , T. secunda sp.n. and T. tertia sp.n. (from La Palma del Condado, Huelva, Spain) are syntopic, and T. quarta sp.n. (from Barrancos, Beja, Portugal) is the first species of the group with umbilicate series 4 + 4. They represent, respectively, the southernmost and westernmost distribution data for the group. The baetica group remains well defined and includes ten species. The diagnosis is simplified, with the confirmation that many morphological features characteristic of this group are widespread within the genus. The defining character is the presence of denticles in apical margin of elytra, not associated to the seventh stria or to the elytral suture. The new species provide morphological data that suggest the baetica and silvanoides groups are closely related. Distribution of both groups in the south of the Iberian Peninsula and the presence of morphological characters that are considered plesiomorphic are coherent with the hypothesis of betic‐riffain origin of the genus. They also support the hypothesis of easy alteration of the umbilicate pattern between closely related species and the independent origin of apical denticles in the group compared with those present in other species of the genus. The presence of three well‐differentiated size ranges in syntopic populations suggest ecological diversification to avoid interspecific competition.     

Genetic analysis based on plastidial and ribosomal sequences of the endemic bi-edaphic taxon Jurinea pinnata (Lag.) DC. (Compositae) in the Guadix-Baza Basin

Jurinea pinnata (Lag.) DC. is one of the three species of the genus that grows in the Iberian Peninsula. This species grows on gypsum and dolomite, substrates rich in endemic taxa. Actually, the genus Jurinea Cass. is particularly rich in endemic taxa associated with special substrates. The island-like distribution of J. pinnata makes it an interesting case for the study of genetic differentiation processes. The geographical proximity of J. pinnata and J. humilis makes this evolutionary puzzle even more difficult to solve. These facts could affect the genetic attributes of the species as far as diversity and differentiation are concerned. Our field research involved the sampling of individuals from eight populations of J. pinnata. Of them, six were located in the Guadix-Baza Basin. We used sequences of ribosomal and plastid DNA to perform the genetic analyses. Results revealed no differences between individuals occurring on the two kinds of substrates and evidences of hybridization between J. pinnata and J. humilis. Furthermore, the phylogenetic analysis revealed a possible polyphyletic origin for the adaptation to special substrates within the Jurinea genus, while monophyly was observed in species from the Iberian Peninsula. This could indicate a more general adaptation to arid or saline environments.     

Pollen morphology of Xanthium L. (s.l.) (Asteraceae,Asteroideae, Heliantheae,Ambrosiinae) in the Iberian Peninsula - a palynotaxonomic approach to a poisonous,allergenic and invasive genus

Abstract

The pollen morphology of the Iberian taxa of the genus Xanthium s.l. and of the non-Iberian species X. ambrosioides was studied to help to identify these poisonous, allergenic and invasive taxa and clarify their taxonomic relationships. In addition, the question of the existence and structure of a mesoaperture in the apertural system of Xanthium was also considered. The pollen grains of sections Acanthoxanthium and Xanthium clearly differ by their characters, which supports the separation of both taxa and allows for their distinction based on pollen morphology. It is suggested that the morphological and palynological differences between Xanthium and Acanthoxanthium, together with the lack of hybridisation between the two sections and the monophyly of Acanthoxanthium, could lead to the elevation of the latter to the generic status. The two species of section Xanthium present in the Iberian Peninsula (X. orientale and X. strumarium) and of section Acanthoxanthium are well separated by their palynological features. Likewise, the Iberian subspecies of Xanthium, X. orientale subsp. orientale and X. orientale subsp. italicum are palynologically distinguishable. However, no significant difference could be found between the pollen of X. strumarium subsp. strumarium and of X. strumarium subsp. brasilicum. The pollen data do not support the hypotheses of a hybrid origin for X. strumarium subsp. brasilicum. A mesoaperture intersecting the foot layer and the upper part of the endexine is present in both sections, which is congruent with the results already found for other Asteroideae.     

Morphometric study of the complex Moehringia sect. Pseudomoehringia McNeill from the western Mediterranean

Moehringia. sect. Pseudomoehringia McNeill is endemic to the western Mediterranean, having its origin and diversification centre in the Iberian Peninsula and mountains of northern Morocco. Both the relationship with the genus Arenaria as well as the taxonomy within the section have been largely controversial. To disentangle these issues, we conducted a morphometric study using herbarium material. We measured 12 relevant morphological traits from 148 selected herbarium sheets. Data were analyzed using different statistical methods: general linear models, multiple factorial analysis and linear discriminant analysis. We found significant differences between Arenaria balearica and the rest of the genus Moehringia taxa. Within this genus, we identified three well-discriminated species: Moehringia fontqueri, M. glochidisperma and M. intricata. Within M. intricata complex, we discerned virtually no differences amongst most of the subspecies (intricata, giennensis and tejedensis) with the exception of subsp. castellana. We propose: (i) to maintain A. balearica separate from Moehringia sect. Pseudomoehringia; (ii) to consider three species in the section Pseudomoehringia: M. fontqueri, M. glochidisperma and M. intricata; (iii) to recognize only two subspecies within M. intricata complex: subsp. intricata and subsp. castellana. To clarify the taxonomy of this threatened group is of great interest because it might help to prioritize conservation measures.     

Appearances can be deceptive: different diversification patterns within a group of Mediterranean earthworms (Oligochaeta, Hormogastridae)

Many recent studies on invertebrates have shown how morphology not always captures the true diversity of taxa, with cryptic speciation often being discussed in this context. Here, we show how diversification patterns can be very different in two clades of closely related earthworms in the genus Hormogaster stressing the risk of using nonspecific substitution rate values across taxa. On the one hand, the Hormogaster elisae species complex, endemic to the central Iberian Peninsula, shows morphological stasis. On the other hand, a clade of Hormogaster from the NE Iberian Peninsula shows an enormous morphological variability, with 15 described morphospecies. The H. elisae complex, however, evolves faster genetically, and this could be explained by the harsher environmental conditions to which it is confined—as detected in this study, that is, sandier and slightly poorer soils with lower pH values than those of the other species in the family. These extreme conditions could be at the same time limiting morphological evolution and thus be responsible for the observed morphological stasis in this clade. Contrarily, Hormogaster species from the NE Iberian Peninsula, although still inhabiting harsher milieu than other earthworm groups, have had the opportunity to evolve into a greater morphological disparity. An attempt to delimit species within this group following the recently proposed general mixed Yule‐coalescent method showed a higher number of entities than expected under the morphospecies concept, most probably due to the low vagility of these animals, which considerably limits gene flow between distant conspecific populations, but also because of the decoupling between morphological and genetic evolution in the H. elisae complex.     

Hypomethylation of the aquatic invasive plant,Ludwigia grandiflora subsp. hexapetala mimics the adaptive transition into the terrestrial morphotype

Ongoing global changes affect ecosystems and open up new opportunities for biological invasion. The ability of invasive species to rapidly adapt to new environments represents a relevant model for studying short-term adaptation mechanisms. The aquatic invasive plant, Ludwigia grandiflora subsp. hexapetala, is classified as harmful in European rivers. In French wet meadows, this species has shown a rapid transition from aquatic to terrestrial environments with emergence of two distinct morphotypes in 5 years. To understand the heritable mechanisms involved in adjustment to such a new environment, we investigate both genetic and epigenetic as possible sources of flexibility involved in this fast terrestrial transition. We found a low overall genetic differentiation between the two morphotypes arguing against the possibility that terrestrial morphotype emerged from a new adaptive genetic capacity. Artificial hypomethylation was induced on both morphotypes to assess the epigenetic hypothesis. We analyzed global DNA methylation, morphological changes, phytohormones and metabolite profiles of both morphotype responses in both aquatic and terrestrial conditions in shoot and root tissues. Hypomethylation significantly affected morphological variables, phytohormone levels and the amount of some metabolites. The effects of hypomethylation depended on morphotypes, conditions and plant tissues, which highlighted differences among the morphotypes and their plasticity. Using a correlative integrative approach, we showed that hypomethylation of the aquatic morphotype mimicked the characteristics of the terrestrial morphotype. Our data suggest that DNA methylation rather than a new adaptive genetic capacity is playing a key role in L. grandiflora subsp. hexapetala plasticity during its rapid aquatic to terrestrial transition.     

Vicariant versus dispersal processes in the settlement of Pseudamnicola (Caenogastropoda,Hydrobiidae) in the Mediterranean Balearic Islands

The genus Pseudamnicola comprises a group of tiny springsnails inhabiting several continental and insular regions of the Mediterranean basin. Given the limited dispersal capabilities of these animals, it is difficult to explain the wide distribution range of the genus and, more specifically, its presence in isolated habitats, such as on islands. Thus, to investigate the process(es) that may explain these distribution patterns, we morphologically re‐described and genetically analysed the six Pseudamnicola (Pseudamnicola) species occurring in the Iberian Peninsula and the nearby Balearic Islands. Genetic relationships were explored by sequencing two mitochondrial (cytochrome c oxidase subunit I and 16S rRNA) and one nuclear (28S rRNA) gene in 19 populations. Our morphological study confirmed the presence of previously described species, whereas our phylogenetic results revealed three lineages within the subgenus: one clade grouping the species from Minorca Island with an Iberian Peninsula species, a second clade grouping the three species from Majorca Island, and a third clade that consists of a single species, which occurs in both the Iberian Peninsula and Ibiza Island. Calculated speciation times show that the cladogenetic events involving the insular species seem to have occurred after the current conformation of the Balearic Islands (c. 20 Mya). Therefore, the speciation process may have been related to subsequent transmarine colonizations, probably during the Messinian Salinity Crisis, and the Pleistocene glaciations when landmass corridors connected the islands with the continent. © 2014 The Linnean Society of London     

Flow cytometric and karyological analyses of Calendula species from Iberian Peninsula

Calendula L. (Asteraceae) is a taxonomically and cytologically complex genus due to its high morphological and karyological variation. To gather consistent cytological information aiming to consolidate the existing knowledge, sustain the taxonomic revision of the genus and explore the evolutionary relationships among species, the genome size and chromosome number of the Iberian Peninsula representatives of this genus were assessed. The study included 11 taxa that occur in the Iberian Peninsula, one in Madeira and two from Morocco. Chromosome counts were made using the squash technique in root tips and flower buds, while nuclear DNA contents were assessed using propidium iodide flow cytometry. The following chromosome numbers are reported: 2n = 44 for C. arvensis, 2n = 30 for C. tripterocarpa, and 2n = 32 for the remaining Iberian taxa. The genome size of Calendula species was assessed for the first time and ranged from 1.75 pg/2C in C. maroccana to 5.41 pg/2C in C. arvensis. Within the complex formed by C. incana and C. suffruticosa, a gradient of genome size values was obtained. Intraspecific variation in genome size was detected in some taxa. The obtained genome size values and their variation are discussed in the light of the theories proposed for the speciation of the genus, with events of hybridization, genome duplication and dysploidy being hypothesized to play a major role in the evolution of this genus.     

First identification of Decennatherium Crusafont, 1952 (Mammalia,Ruminantia, Pecora) in the Siwaliks of Pakistan

Previously undescribed remains of a new large giraffid have been identified from the late Miocene Siwaliks Hills of the Potwar Plateau in Pakistan. This taxon is very intimately related to the late Miocene giraffid genus Decennatherium, previously only certainly identified in the Iberian Peninsula, and with some possible remains assigned to Decennatherium crusafonti from Aliäbäd (Iran). The new material collected in the Siwaliks Hills shows a high morphological similarity with the Spanish remains of the genus, especially the early Vallesian Decennatherium pachecoi. Two previously undescribed ossicone fragments from the same area are also described and identified as cf. Decennatherium. These findings represent the easternmost occurrence of the genus Decennatherium and show a late Miocene migration of the genus throughout the Iberian Peninsula and Southern Asia.     

Ecological differentiation,speciation, and rarity: How do they match in Tephroseris longifolia agg. (Asteraceae)?

Tephroseris longifolia agg. is a complex group of outcrossing perennials distributed throughout Central Europe. Recent morphological study revealed six morphotypes corresponding to five previously distinguished subspecies, together with Alpine and Pannonian morphotypes of T. longifolia subsp. longifolia. The delimited morphotypes differ in relative DNA content, geographical range, and rarity. We compared ecological niches of the six morphotypes in order to assess the impact of ecological differentiation on the speciation processes within the T. longifolia agg. Further, we examined whether morphotypes with small range are more ecologically specialized than their widespread relatives. The distribution area of the aggregate includes the Alps, Apennines, Carpathians, and the Pannonian Basin. Ecological variables linked to climate, topography, soil, and vegetation were gathered from 135 circular plots recorded in 35 localities. Related variables were grouped to describe the partial ecological niches: climatic, topographic, pedological, biotic, and coenotic (based either on vascular plants or on bryophytes), each of them visualized as an envelope in the two‐dimensional nonmetric multidimensional scaling ordination space. Each partial ecological niche for a given morphotype was characterized by its position (location of the envelope centroid), breadth (surface of the envelope), and overlaps with envelopes of the other morphotypes. Mantel statistics based on Spearman correlation coefficients were used to quantify differentiation of morphotypes in ecological parameters represented by the partial ecological niches. The significant niche differentiation was confirmed for climatic, topographic, pedological, and vascular plant‐based coenotic niches. Ecological niche differentiation corresponded well to morphological and partially also to karyological differentiation. Narrowly distributed morphotypes occupied more specific habitats and had narrower ecological niches than their widespread relatives. Ecological differentiation could be considered an important driver in allopatric speciation within the T. longifolia agg. Our results demonstrate that quantification of ecological divergence is helpful in assessing evolutionary history of closely related taxa.